Dietary composition and diel feeding patterns of epipelagic siphonophores

Prey consumption patterns are described for 24 species of epipelagic siphonophores studied during 1977–1980 in the Gulf of California, off Southern California, in the Sargasso Sea, and in Friday Harbor, Washington. Of the species, 7 were studied by day and at night, 15 were studied only by day, and 2 were studied only at night. Each of the 3 suborders of siphonophores had a characteristic diet. Siphonophores in the suborder Cystonectae, which had large gastrozooids, fed primarily on fish larvae. Species in the suborder Physonectae, which generally had few, large gastrozooids, consumed some small copepods, but consumed mainly large copepods and a variety of large, noncopepod prey. Species in the suborder Calycophorae, which generally had many small gastrozooids, fed mainly on small copepods, and also on other small prey organisms. The maximum size of prey tended to be correlated with gastrozooid length for all the siphonophores studied. For a given siphonophore species, the number of ingested prey was greatest at localities where prey organisms were most abundant in the surrounding seawater. For siphonophore species collected both day and night, there was a tendency for more prey to be consumed at night. Behavioral observations in the laboratory indicated that of 7 siphonophore species tested, 2 fed only in the light, and another fed only in the dark.     

Metabolic rates of epipelagic marine zooplankton as a function of body mass and temperature

The metabolic rates (oxygen uptake, ammonia excretion, phosphate excretion) of epipelagic marine zooplankton have been expressed as a function of body mass (dry, carbon, nitrogen and phosphorus weights) and habitat temperature, using the multiple-regression method. Zooplankton data used for this analysis are from phylogenetically mixed groups (56 to 143 species, representing 7 to 8 phyla, body mass range: 6 orders of magnitude) from various latitudes (habitat temperature range:-1.4° to 30°C). The results revealed that 84 to 96% of variation in metabolic rates is due to body mass and habitat temperature. Among the various body-mass units, the best correlation was provided by carbon and nitrogen units for all three metabolic rates. Oxygen uptake, ammonia excretion and phosphate excretion are all similar in terms of body-mass effect, but differ in terms of temperature effect. With carbon or nitrogen body-mass units, calculated Q₁₀ values are 1.82 to 1.89 for oxygen uptake, 1.91 to 1.93 for ammonia excretion and 1.55 for phosphate excretion. The effects of body mass and habitat temperature on the metabolic quotients (O:N, N:P, O:P) are insignificant. The present results for oxygen-uptake rate vs body mass do not differ significantly from those reported for general poikilotherms by Hemmingsen and for crustaceans by Ivleva at a comparable temperature (20°C). The importance of a body-mass measure for meaningful comparison is suggested by the evaluation of the habitat-temperature effect between mixed taxonomic groups and selected ones. Considering the dominant effects of body mass and temperature on zooplankton metabolic rates, the latitudinal gradient of community metabolic rate for net zooplankton in the ocean is estimated, emphasizing the non-parallelism between community metabolic rates and the standing stock of net zooplankton.     

Distribution and trophic links of gelatinous zooplankton on Dogger Bank,North Sea

The ecology of small, gelatinous zooplankton is not integrated into management of Dogger Bank (54° 00′ N, 3° 25′ E to 55° 35′ N, 2° 20′ E). In pursuit of this goal, gelatinous zooplankton and their potential prey were sampled along a transect across the bank on June 10–16, 2007. Eleven species of small medusae and ctenophores were collected, with six abundant taxa occurring in greater numbers below the thermocline and in the shallower, southeastern portion of the bank. There were no statistically significant diel changes in distribution. In contrast, potential prey were distributed more evenly across the bank and throughout the water column. Isotopic analyses revealed that gelatinous zooplankton fed on both smaller (100–300 μm) and larger (>300 μm) mesozooplankton, but also potentially on each other. These ecological insights suggest that small medusae and ctenophores should be integrated into sustainable management of Dogger Bank.     

Estimating zooplankton biomass through image analysis

Zooplankton biomass can be reliably estimated by a non-destructive method based on the statistical relationship between biovolume and the contents of carbon and nitrogen. Integrated zooplankton biovolume and the corresponding organic C and N contents were analysed in paired fresh and fixed (hexamine-buffered formalin) samples. Sample biovolume was estimated through the integration of individual volumes obtained by semi-automatic image analysis on fixed zooplankton samples. The corresponding C and N contents of paired fresh and preserved zooplankton were analysed with a Carlo-Erba C-H-N Analyser. The method allows simultaneous measurement of integrated and individual numbers and biomass (biovolume, C and N) spectrum. It is non-destructive and allows the minimization of errors and variability attributable to differences in the handling processes of samples and to the presence of particles other than zooplankton, like phytoplankton or detritus particulate matter.     

Bioluminescence spectra of shallow and deep-sea gelatinous zooplankton: ctenophores, medusae and siphonophores

We have examined the variability and potential adaptive significance of the wavelengths of light produced by gelatinous zooplankton. Bioluminescence spectra were measured from 100 species of planktonic cnidarians and ctenophores collected between 1 and 3500 m depth. Species averages of maximal wavelengths for all groups ranged from 440 to 506 nm. Ctenophores (41 species) had characteristically longer wavelengths than medusae (34 species), and the wavelengths from siphonophores (25 species) had a bimodal distribution across species. Four species each produced two different wavelengths of light, and in the siphonophore Abylopsistetragona these differences were associated with specific body regions. Light from deep-dwelling species had significantly shorter wavelengths than light from shallow species in both ctenophores (p = 0.010) and medusae (p = 0.009). Although light production in these organisms was limited to the blue-green wavelengths, it appears that within this range, colors are well-adapted to the particular environment which the species inhabit. Received: 27 April 1998 / Accepted: 27 October 1998     

Sinking rates of fecal pellets from gelatinous zooplankton (Salps,Pteropods, Doliolids)

Sinking rates were determined for fecal pellets produced by gelatinous zooplankton (salps, Salpa fusiformis and Pegea socia; pteropods, Corolla spectabilis; and doliolids, Dolioletta gegenbaurii) feeding in surface waters of the California Current. Pellets from the salps and pteropods sank at rates up to 2 700 and 1 800 m d^-1, respectively; such speeds exceed any yet recorded for zooplankton fecal pellets. Fecal pellets of salps were rich in organic material, with C:N ratios from 5.4 to 6.2, close to values for living plankton. The relation between volume and sinking rate indicates that salp and pteropod pellets are slightly less dense than those of pelagic Crustacea; moreover, pellet density varied between different collection dates, probably because of differences in composition. In contrast, doliolid pellets sank at rates up to 208 m d^-1, a rate much lower than would be expected from pellet size. Thus, density and sinking rates of pellets are much more variable in zooplankton than would be expected from studies of crustaceans alone. Moreover, the extraordinarily high sinking rates of fecal pellets of salps indicates that these tunicates may be disproportionately important in the flux of biogenic materials during periods when they form dense population blooms.     

Preservation methods alter stable isotope values in gelatinous zooplankton: implications for interpreting trophic ecology

Jellyfish are increasingly topical within studies of marine food webs. Stable isotope analysis represents a valuable technique to unravel the complex trophic role of these long-overlooked species. In other taxa, sample preservation has been shown to alter the isotopic values of species under consideration, potentially leading to misinterpretation of trophic ecology. To identify potential preservation effects in jellyfish, we collected Aurelia aurita from Strangford Lough (54^o22′44.73″N, 5^o32′53.44″W) during May 2009 and processed them using three different methods prior to isotopic analysis (unpreserved, frozen and preserved in ethanol). A distinct preservation effect was found on δ¹⁵N values: furthermore, preservation also influenced the positive allometric relationship between individual size and δ¹⁵N values. Conversely, δ¹³C values remained consistent between the three preservation methods, conflicting with previous findings for other invertebrate, fish and mammalian species. These findings have implications for incorporation of jellyfish into marine food webs and remote sampling regimes where preservation of samples is unavoidable.     

A comparison of carbon based ammonia excretion rates between gelatinous and non-gelatinous zooplankton: Implications and consequences

About 560 literature data on weight-specific ammonia excretion rates of gelatinous zooplankton (cnidarians, ctenophores and salps) and non-gelatinous zooplankton (mainly crustacea) were converted to carbon based units to enabel a better comparison between both groups. If carbon is used as body-mass unit ammonia excretion rates of gelatinous zooplankton are in the same range as values obtained for other zooplankton taxa, indicating a similar nitrogen output per unit of organic matter in both groups. These results suggest nutrient regeneration potential to be the same in gelatinous and non-gelatinous zooplankton, and that nutrient regeneration within the pelagic system depends more upon the carbon biomass ratio between gelatinous and other zooplankton than on physiological differences.     

Prediction of response of zooplankton biomass to climatic and oceanic changes

This paper examines the long-term variation in zooplankton biomass in response to climatic and oceanic changes, using a neural network as a nonlinear multivariate analysis method. Zooplankton data collected from 1951 to 1990 off the shore of northeastern Japan were analyzed. We considered patterns of the Kuroshio and the Oyashio, sea surface temperature, and meteorological parameters as environmental factors that affect zooplankton biomass. Back propagation neural networks were trained to generate mapping functions between environmental variables and zooplankton biomass. The performance of the network models was tested by varying the numbers of input and hidden units. Changes in zooplankton biomass could be predicted from environmental conditions. The neural network yielded predictions with smaller errors than those of predictions determined by linear multiple regression. The sensitivity analysis of networks was used to extract predictive knowledge. The air pressure, sea surface temperature, and some indices of atmospheric circulation were the primary factors for predictions. The patterns of the Kuroshio and the Oyashio demonstrated different effects among sea areas.     

Relation of fish larvae and zooplankton biomass in the Gulf of Aden

This study is based on zooplankton samples collected in the upper 50 m by the F.R.V. Manihine in the Gulf of Aden during October–November, 1966 and February–March, 1967. Generally, the displacement volume of zooplankton varied between 20 and 67 ml/m². Some higher values, up to 100 ml/m², were also observed. The number of larval fish in positive hauls ranged from 2 to 282 larvae/m². An inverse relationship between the number of larval fish and the accompanying volume of zooplankton was noted. This relationship is discussed. It is hypothesized that larval mortality due to predation in the Gulf of Aden during the northeast monsoon (November, February and March) was very high.     

Nutrient regeneration by zooplankton during a red tide off Peru,with notes on biomass and species composition of zooplankton

During March and April 1976, a red tide, dominated by the dinoflagellate Gymnodinium splendens Lebour, developed in the vicinity of 15°06'S and 75°31'W off Peru. At the height of the bloom, the euphotic zone was 6 m deep and the chlorophyll a at the surface was 48 g l^-1. A daily collection of zooplankton at 09.00 hrs showed large fluctuations of biomass, from 0.2 to 3.84 g dry weight m^-2 in a water column of 120m. Copepodids and nauplii dominated the collections. During a period of reduced wind, the adult copepods were a mixture of the species characteristic of the coastal upwelling system and the neritic species associated with more northerly, tropical waters. Nitrogen regeneration by the zooplankton varied with the development of the bloom, the type of zooplankton dominating the experiment, and biomass fluctuations, but never accounted for more than 25% of the nitrogen uptake by phytoplankton.     

Metabolism of epipelagic tropical ctenophores

Measurements of respiration and excretion at 25°C were made for five species of ctenophores collected during five cruises to the Bahamas (1982–1984). The mean element-specific respiration and ammonium excretion rates of freshly collected specimens of all species ranged from 4 to 16% d^-1, the mean atomic O:N ratios were 10 to 16, and ammonium averaged 60 to 90% of the total dissolved nitrogen excreted. For adult ctenophores, the carbon content ranged from 0.6% carbon (as percent of dry weight) for Bolinopsis vitrea to 3.7% carbon for Beroë ovata. There was a marked increase in the organic content (% carbon of dry weight) of small Bolinopsis vitrea with tentacles compared to fully lobate adults. B. vitrea had increasingly higher metabolic rates when held at food concentrations up to 100 copepods 1^-1 (about 250 g C 1^-1). The overall range between starved and well-fed B. vitrea was about two times for respiration and a factor of three for ammonium excretion. B. vitrea decreased from well-fed to a starved metabolic rate in about a day after removal from food. The metabolic rate of Eurhamphaea vexilligera was not measurably affected by short-term starvation or feeding (maximum 25 copepods 1^-1). In feeding experiments, E. vexilligera of 20 to 56 mm length fed at rates equivalent to clearance rates of 250 to 1 800 ml h^-1.     

Global biodiversity patterns of benthic marine algae

Species richness patterns are remarkably similar across many marine taxa, yet explanations of how such patterns are generated and maintained are conflicting. I use published occurrence data to identify previously masked latitudinal and longitudinal diversity gradients for all genera of benthic marine macroalgae and for species in the Order Bryopsidales. I also quantify the size, location, and overlap of macroalgal geographic ranges to determine how the observed richness patterns are generated. Algal genera exhibit an inverse latitudinal gradient, with biodiversity hotspots in temperate regions, while bryopsidalean species reach peak diversity in the tropics. The geographic distribution of range locations results in distinct clusters of range mid-points. In particular, widespread taxa are centered within tight latitudinal and longitudinal bands in the middle of the Indo-Pacific and Atlantic Oceans while small-ranged taxa are clustered in peripheral locations, suggesting that variation in speciation and extinction are important drivers of algal diversity patterns. Hypotheses about factors that regulate diversity contain underlying assumptions about the size and location of geographic ranges, in addition to predictions as to why species numbers will differ among regions. Yet these assumptions are rarely considered in assessing the validity of the prevailing hypotheses. I assess a suite of hypotheses, suggested to explain patterns of marine diversity, by comparing algal-richness patterns in combination with the size and location of algal geographic ranges, to the richness and range locations predicted by these hypotheses. In particular, the results implicate habitat areas and ocean currents as the most plausible drivers of observed diversity patterns.     

Above and below ground biomass patterns in arid lands

Regular vegetation patterns have been observed in many arid zones around the word. This particular spontaneous arrangement of the vegetation optimizes the use of the scarce water resources and could be imitated to restore vulnerable ecosystems; at the same time, the patterns of vegetation act as an early warning signal of the fact that fragile ecosystems may suddenly undergo irreversible shifts, thus, they deserve a special attention. The formation of vegetation patterns is the object of many theoretical and experimental studies, nevertheless, in previous works, the interest that is deserved to below ground biomass allocation is minor as compared to the effort that is spent to describe the organization of vegetation above ground.     

Temporal, spatial, and between-host comparisons of patterns of parasitism in lake zooplankton

In nature, multiple parasite species infect multiple host species and are influenced by processes operating across different spatial and temporal scales. Data sets incorporating these complexities offer exciting opportunities to examine factors that shape epidemics. We present a method using generalized linear mixed models in a multilevel modeling framework to analyze patterns of variances and correlations in binomially distributed prevalence data. We then apply it to a multi-lake, multiyear data set involving two Daphnia host species and nine microparasite species. We found that the largest source of variation in parasite prevalence was the species identities of host-parasite pairs, indicating strong host-parasite specificity. Within host-parasite combinations, spatial variation (among lakes) exceeded interannual variation. This suggests that factors promoting differences among lakes (e.g., habitat characteristics and species interactions) better explain variation in peak infection prevalence in our data set than factors driving differences among years (e.g., climate). Prevalences of parasites in D. dentifera were more positively correlated than those for D. pulicaria, suggesting that similar factors influenced epidemic size among parasites in D. dentifera. Overall, this study demonstrates a method for parsing patterns of variation and covariation in infection prevalence data, providing greater insight into the relative importance of different underlying drivers of parasitism.     

A comparison of zooplankton patterns in the California Current and North Pacific Central Gyre

This study compares the small-scale (100's to 1000's of meters) horizontal spatial distribution of zooplankton from stations in the California Current (29°N; 118°W) and in the North Pacific Central Gyre (31°N; 155°W). Patterns were inferred from the spacing of vertical tows and the variability in abundance of 15 taxonomic categories caught in these tows. In the California Current, 4 tows were taken at random positions within 2000 m of a drogue or fixed geographic position and 4 replicate tows were taken at the drougue or the fixed point. Four series of these 8 tows were taken around noon and midnight during two days. At the Central Gyre station, two similar series (one day and one night) were taken while following a drogue. Estimates of the scale of structure based on comparisons of replicate and random tow variability suggest that aggregated patterns in the Central Gyre are larger than a few hundred meters, while the California Current structures may be less than 100 m in size. Day-night changes in patch size were not apparent in either area. Evidence for multispecies patches was found only for the Central Gyre station. Replication samples in both areas gave more consistent measures of community structure (relative proportions of species) than the random tows. Similarity of community structure decreased with increasing distance between tows being compared. The decrease was much greater for the California Current station, a further indication of smaller patch size in this area. Diurnal changes in community structure occurred in both areas.Based in part on a dissertation submitted in partial fulfillment of the requirements of the Ph.D. degree at the University of California, San Diego. Portions of this research were supported by the Marine Life Research Program of the Scripps Institution of Oceanography.     

Diurnal rhythm of the bioluminescent field in the ocean epipelagic zone

Characteristics of the diurnal rhythm of the bioluminescence field are analyzed on the basis of experimental data obtained in the region around Kamchatka peninsula (50°31′N; 163°50′E). The feasibility of reconstructing the nocturnal structure of the bioluminescence field by measurements made at other times of the day has been demonstrated. Diurnal rhythm parameters for the bioluminescent field of several other studied areas of the ocean have been defined. The temporal parameters feature linear latitudinal dependence. At the equator the boundaries of morning and evening transition periods are 05:06 to 07:30 and 17:48 to 19:54 hrs, respectively. In the Barents Sea in spring these boundaries are 02:42 to 04:06 and 19:48 to 21:12 hrs. Boundary times of diurnal rhythm do not deviate from the linear regression by more than 10%. Received: 21 August 1998 / Accepted 4 March 1999     

Spectral composition of bioluminescence of epipelagic organisms from the Sargasso Sea

The spectral characteristics of single identified epipelagic sources of bioluminescence from the western Sargasso Sea were measured with an optical multichannel analyzer (OMA) system during the April, 1985, Biowatt cruise. The emission spectra of specimens representing 45 species from 8 phyla were measured. Peak bioluminescence emissions typically occurred between 440 and 500 nm, in the blue region of the visible spectrum. Three exceptions involved emission in the green, yellow, and red spectral regions. Intraspectific variability in spectra, was noted in several species. One shrimp species exhibited two modes of light emission, each with different emission spectra. Other cases involved dynamic color shifts of 10 to 14 nm; the source of the spectral variability is unknown, but may involve optical filtering or differences in the color of luminescence from multiple sites of light emission. Measurements from independent samples of unsorted plankton revealed different spectral distributions. This suggests that the spectral emissions of bioluminescence in the upper water column will vary, based on species assemblage.     

Effects of a decrease in downwelling irradiance on the daytime vertical distribution patterns of zooplankton and micronekton

The daytime vertical distribution of several species of crustaceans, gelatinous zooplankton, and fish were monitored in situ simultaneously with measurements of downwelling irradiance in Oceanographer Canyon in July 1999. During this submersible-based research cruise, an influx of turbid water significantly decreased downwelling irradiance and had a substantial impact on the depth distributions of a number of organisms. Several species of crustaceans, (Thysanoessa gregaria and Sergestes arcticus) and gelatinous zooplankton (Salpa aspera and Salpa fusiformis) ascended over 100 m in the water column during the influx and returned to their pre-influx depths once the influx had ceased. In situ light measurements demonstrated that each of these species was associated with the same irradiance levels during the influx as they were under pre- and post-influx conditions. By contrast, a statistically significant change in temperature, salinity, and oxygen concentrations measured post-influx had no apparent impact on the depth distributions. These results indicate that these species were adjusting their depth distributions to remain within a range of preferred irradiances. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s002270020788